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M590p155.Pdf Vol. 590: 155–169, 2018 MARINE ECOLOGY PROGRESS SERIES Published March 12 https://doi.org/10.3354/meps12480 Mar Ecol Prog Ser OPENPEN ACCESSCCESS Constraining species−size class variability in rates of parrotfish bioerosion on Maldivian coral reefs: implications for regional-scale bioerosion estimates Robert T. Yarlett1,*, Chris T. Perry1, Rod W. Wilson2, Kate E. Philpot3 1Geography, College of Life and Environmental Sciences, University of Exeter, Exeter EX4 4RJ, UK 2Biosciences, College of Life and Environmental Sciences, University of Exeter, Exeter EX4 4QD, UK 3Ecology by Design Ltd, Unit 16, Hampden House, Monument Park, Chalgrove, Oxfordshire OX44 7RW, UK ABSTRACT: Parrotfish are important bioeroders on coral reefs, and thus influence reef carbonate budgets and generate large volumes of carbonate sand that contribute to local beach and reef island maintenance. However, despite the importance of this process, there is a paucity of data with which variations in bioerosion rates as a function of species, feeding modes, and body size of parrotfish can be constrained. There is, in addition, limited knowledge regarding how resultant rates may vary within and between reef-building regions. Here, direct estimates of parrotfish bio- erosion rates were quantified across different size classes of 6 common species of Maldivian par- rotfish. These species comprise both ‘scraper’ and ‘excavator’ taxa, and our data indicate marked variations in mean bioerosion rates among these species. We also note that all species exhibited an apparent bimodal feeding cycle, with peaks in the late morning and early afternoon. Highest bioerosion rates were found in the ‘excavator’ Chlorurus strongylocephalus (~460 kg ind.−1 yr−1), nearly 130 times greater than rates calculated for comparably sized (>45 cm) ‘scraper’ species. Our data provide metrics that can be used in conjunction with parrotfish biomass or density data to improve estimates of parrotfish bioerosion on central Indian Ocean reefs, a region of high par- rotfish density, but from which only limited metrics exist. We emphasise the importance of obtain- ing sub-regional scale process data to better inform estimates of reef bioerosion, especially to sup- port attempts to model the impacts of fishing pressure, which commonly results in removal of high-rate bioeroding taxa. KEY WORDS: Parrotfish · Bioerosion · Maldives · Coral reefs INTRODUCTION shaped by a combination of biological, physical, and chemical constructional and erosional processes The structural complexity and growth potential of (Scoffin 1992, Perry & Hepburn 2008). Framework coral reefs underpin many reef ecosystem services, construction is primarily the result of the production such as shoreline protection, and habitat provision of carbonate skeletons by corals, while the most per- for a diverse array of marine organisms, including vasive form of erosion is that by bioeroding organ- many commercially important species (Lugo-Fernán- isms. This leads to weakening or erosion of the reef dez et al. 1998, Moberg & Folke 1999, Ruckelshaus et substrate (Stearn et al. 1977, Scoffin et al. 1980, al. 2013, Ferrario et al. 2014). These systems are Glynn 1997). Assessments of rates of carbonate accu- © The authors 2018. Open Access under Creative Commons by *Corresponding author: [email protected] Attribution Licence. Use, distribution and reproduction are un - restricted. Authors and original publication must be credited. Publisher: Inter-Research · www.int-res.com 156 Mar Ecol Prog Ser 590: 155–169, 2018 mulation (e.g. by corals and coralline algae, and by (1996) in the Caribbean. While more recent studies sediment producers such as Halimeda spp. and Fora - have quantified these processes for more species in minifera), less that lost through bioerosion (e.g. by different regions (e.g. Great Barrier Reef: Bellwood fish, urchins, sponges, and microborers) can thus be et al. 2003; Red Sea: Alwany et al. 2009; Hawaii: used to measure reef carbonate budgets (sensu Perry Ong & Holland 2010), our understanding of the et al. 2008), which can provide an indication of net variability in these processes between species, sizes, reef framework accumulation or loss. As a result, car- and geographic locations remains very limited. bonate budget assessments are becoming increas- There are ~99 recognised species of parrotfish world- ingly relevant in the light of recent global coral wide, and >70 species are categorised into scraping bleaching events, which have caused large-scale or excavating feeding modes, many of which are coral mortality in a number of regions (Hughes et al. geographically widespread (Choat et al. 2012). Yet 2017), and with potential negative impacts on reef direct estimates of parrotfish bioerosion rates are growth capacity (Perry & Morgan 2017). restricted to data from just 15 species from specific On the erosional side of the carbonate budget locations (Bellwood 1995a, Bruggemann et al. 1996, question, parrotfish (family Labridae) are often iden- Bellwood et al. 2003, Alwany et al. 2009, Ong & tified as especially important bioeroders (e.g. Bell- Holland 2010, Morgan & Kench 2016). There is wood et al. 2003, Perry et al. 2015a). Whilst feeding therefore little understanding of how bioerosion primarily on dead coral and rubble substrates rates vary both among species, and among closely (Bruggemann et al. 1994a, Bellwood 1995b, Afeworki related species in different regions, with much of et al. 2011), many parrotfish take bites out of the reef the current data restricted to the largest terminal- framework, likely targeting cyanobacteria (Clements phase males (see Bruggemann et al. 1996, Ong & et al. 2016). This framework material is ingested Holland 2010 for exceptions). In addition, studies along with organic matter, broken down by modified examining how bioerosion rates differ be tween gill arch elements known as the pharyngeal mill scraping and excavating species are sometimes (Bellwood & Choat 1990, Carr et al. 2006), processed contradictory (Bruggemann et al. 1996, Alwany et in the gut, and egested as sediment (Bellwood 1995b, al. 2009, Ong & Holland 2010). Some further studies 1996, Morgan & Kench 2016). These parrotfish can contribute useful data on parrotfish bite rates and be categorised into ‘scraping’ or ‘excavating’ feeding grazing scar dynamics in the context of algal modes, which are defined based on their musculo- grazing (such as Fox & Bellwood 2007, Bonaldo & skeletal systems around the jaw, and feeding behav- Bellwood 2008, Lokrantz et al. 2008, Bejarano et al. iour (Bellwood & Choat 1990). These bioerosion and 2013); however, even with this additional data, sediment-generation processes are increasingly re- accessible datasets on parrotfish bioerosion rates cognised not only as an important component in coral are limited, given their diversity and geographic reef carbonate budgets (Perry et al. 2014), but also distribution. as an important source of sediment to both reef and The present study aimed to address a key geo- lagoonal sediments (Scoffin et al. 1980), and to graphic gap with respect to parrotfish bioerosion reef-associated landforms such as reef islands and data by presenting rates, as well as associated feed- beaches (Perry et al. 2015b, 2017, Morgan & Kench ing metrics, for 6 of the most common species present 2016). on Maldivian coral reefs. The central In dian Ocean Much of the current work on parrotfish functional remains an area where parrotfish populations are roles is summarised in Bonaldo et al. (2014), but reported to be relatively healthy compared to other early work by Gygi (1975), Ogden (1977), Frydl & regions due to the lack of reef-based fishing pressure Stearn (1978), and Scoffin et al. (1980) in the Carib- (McClanahan 2011) and are the most important bio- bean highlighted the importance of parrotfish bio- eroding organisms in the region as a result (Perry et erosion. These early studies used estimates of daily al. 2015b, 2017), yet region-specific rates are limited gut throughput and sediment content in the gut to (but see Morgan & Kench 2016 for data on 2 species). estimate bioerosion and sediment reworking rates The species studied are representative of the range (the ingestion, processing, and egestion of loose of sizes (including both initial and terminal life sediment on the reef). More direct estimates of par- phases) and feeding modes of parrotfish found in the rotfish bioerosion, involving observations of daily region. In addition, we consider how the bioerosion feeding rates and measures of grazing scar dimen- rate estimates from the present study compare with sions, were then introduced by Bellwood (1995a) on published data on the same, or closely related spe- the Great Barrier Reef and Bruggemann et al. cies in different regions; work that highlights the Yarlett et al.: Parrotfish bioerosion in the Maldives 157 importance of collecting local rate data to inform bio- Species selection erosion estimates. At the primary study site (Vavvaru Island), 15 spe- cies of parrotfish across 5 genera (Chlorurus spp., MATERIALS AND METHODS Scarus spp., Cetoscarus spp., Hipposcarus spp., and Calatomus spp.) were identified from a range of reef Study area habitats across the reef platform. Based on prelimi- nary measures of numerical dominance (Perry et al. Data were collected during field seasons in early 2017), and to ensure representation of the full range 2015 and 2016, primarily from an atoll edge reef plat- of sizes (including both initial and terminal life form site (Vavvaru, Lhaviyani Atoll) in the northern- phases) and feeding modes (both scrapers and exca- central Maldives (Fig. 1; 5° 25’ 5.0’’ N, 73° 21’ 14.0’’ E), vators), the following species were chosen for and augmented with additional data on the same spe- focussed study (species and total length, excluding cies collected in the southern Maldives (Kandahalagala filaments); excavators: Chlo rurus sordidus (up to and Maahutigalaa, Gaafu Dhaalu Atoll), also during ~40 cm) and C. strongylocephalus (up to 70 cm, but the present study in early 2016. The Maldives experi- few over ~60 cm), and scrapers: Scarus frenatus (up ences 2 monsoon periods, with winds from the west- to ~50 cm), S. niger (up to ~45 cm), S.
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